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('Mitrastemoneae'). Maki- No's Initial Use in 1909 of This Name (As 'Mitrastemmaeae') Is Invalid As No Descrip- Tion in Either Japanese Or English Was Given (Dr

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('Mitrastemoneae'). Maki- No's Initial Use in 1909 of This Name (As 'Mitrastemmaeae') Is Invalid As No Descrip- Tion in Either Japanese Or English Was Given (Dr BLUMEA 38 (1993) 221-229 A revision of Mitrastema (Rafflesiaceae) W. Meijer & J.F. Veldkamp Summary Makino called Mitrastemon has Mitrastema (Rafflesiaceae), usually incorrectly Makino, two spe- cies, one in Southeast Asia and one in Central and South America. Introduction Mitrastema was described by Makino (1909) on M. (‘Mitrastemma’) yamamotoi Makino. He intended to name the genus after the mitre-shaped staminal tube, as becomes evident when in 1911 he changed the name to Mitrastemon.This change is On the other Stearn not allowed and Mitrastemon is a superfluous name. hand, as (1973: 519) has pointed out, the Greek words 'sterna' (thread, stamen) and 'stemma' confused, ‘Mitrastemma’ be (garland or wreath) are easily and so may regarded as an orthographic error to be corrected under Art. 73.1 (Exx. 2, 3). ‘Mitrastemon’ or ‘Mitrastemma’ have been adopted in laterpublications, and even by the International Code of Botanical Nomenclature, where the name Mitrastemonaceae, invalid because it is based on an invalid name contrary to Art. 18.1, is conserved (1988: 104). Since its description Mitrastema yamamotoi has become one of the most widely of the Rafflesiaceae. Watanabe's investigated taxa Especially papers on its morphol- and of devoted it is still ogy biology (1933-1937) are masterpieces study; yet not clear what the precise taxonomic position of the genus is and whether it really be- longs to the Rafflesiaceae. Some authors have regarded it as a distinct family, Mitrastemonaceae Makino (1911) (nom. cons.) (e.g. Cronquist, 1981; 'Mitrastemmataceae', Mabberley, 1987), treatedit Rafflesiaceae tribe others have as Mitrastemateae ('Mitrastemoneae'). Maki- no's initial use in 1909 of this name (as 'Mitrastemmaeae') is invalid as no descrip- tion in either Japanese or English was given (Dr. J.N. Westerhoven, Hirosaki, oral validated comm.). It was by Hayata (1913) ('Mitrastemoneae'). Makino it distinct order Mitrastemonales (1911: 252) even proposed to represent a next to the Nepenthales, but he was apparently never followed in this. Although he called it 'series Mitrastemonales' contrary to Art. 33.4 the name seems valid, if one that 'series' here is the latinizationof the German accepts 'Reihe' of Art. 17.2. Cronquist, hedging his bets, wrote “‘Mitrastemon’ has often been included in the Rafflesiaceae, to which it is evidently related. It is more primitive than the Rafflesia- !) School of Biological Sciences, University of Kentucky, Lexington, KY 40506, U.S.A. 2 ) Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands. 222 BLUMEA Vol. 38. No. 1, 1993 ceae in its hypogynous, perfect flowers, and its dome-like synandrium is more near- ly comparable to that of Prosopanche, in the Hydnoraceae, than to anything in the Rafflesiaceae. WhetherMitrastemon should be included in the Rafflesiaceae or kept is of taste." as a separate family largely a matter To us Hayata (1913) has given overwhelming evidence that the genus belongs used the extended Watanabe to Rafflesiaceae. He following arguments, by (1933- 1937) in great detail: characteristics different from from 1) The endophytic body shows no those known the Rafflesiaceaeproper. 2) Bisexual flowers occur in Rhizanthes zippelii (Blume) Spach and Bdallophyton oxylepis (B.L. Rob.) Harms. 3) The tubularandroecium is not completely unique to Mitrastema. In some species the also form tube of the East African Berlinianche (Harms) Vattimo stamens a Vattimo around the pistil, e.g. in B. aethiopica (Welw.) and B. holtzii (Engl.) Vattimo. 4) The superior ovary is an insufficient reason to exclude Mitrastema from the Raf- families well flesiaceae. In many other genera with superior as as inferior ovaries have been admitted. Thus, not surprisingly, semi-inferiorovaries are known in of in blanchetii some species Pilostyles Guill., e.g. P. R.Br. (Gardner, 1844), P. sessilis Rose (1909), P. thurberi A.Gray (Gray, 1854), and P. ulei Solms- Laub. (Harms, 1935), at least in the female flowers. 5) All the detailedstructures of the embryology, ovules, and seeds of Mitrastema agree well with those known for other species of Rafflesiaceae proper. The multibranched, multifloweredinflorescences of Cytinus L. and Bdallophyton Eichl. and that these the closest seem primitive so suggest may represent genera to chlorophyllous ancestors of Rafflesiaceae. All other genera are single-flowered, though especially in Apodanthes Poit. and Pilostyles it might be considered that the inflorescences buried inside the host and that flowers are their pop up through the bark. In Mitrastema the flowers are still very gregarious, but they have become less so in Rafflesia Blume, Rhizanthes Dumort., and Sapria Griff. Mitrastema also shares withother genera of the family the unicellularovary with parietal placentas. Watanabe (1936c) concludedthat the stamen tube is derived from 15-20 (average 16) extrorse anthers, each originally with 4 pollen sacs and a connective extension. It is pushed off after the pollen has been shed by the growth of the pistil, ensuring cross-fertilization (the anthers are on the outside of the tube, whilethe stigma is inside it). Sexual reproduction is apparently obligatory (Watanabe, 1934a) but not always very successful. At higher latitudes the number of fertile fruits rapidly decreases. He mentioned (1933a) that ripe fruits always become infested by a fungus. Its concentrated the chlamydospores are around micropylar area. He suggested (1936d) that the be fungus might fungicidal to other fungi. It may perhaps also take part in germination. It is not clear how many species there actually are. Opinions vary between one to five, or even more. W. Meijer & J.F. Veldkamp: Revision of Mitrastema 223 Makino (1928), Matsuda ('Matuda') (1947), Van Royen (1963), and Hansen (1973) have defended the opinion that there is only one. Corner & Watanabe(1969) depicted what they considered to be 5 species, and Corner (1978) could not identify the population from the Pinosok Plateau, Kinabalu, Sabah, which is now destroyed, so suggesting it was an undescribed one. Watanabe (1934d, 1936a) described geographic variation in Taiwan and Japan in the numberof pairs of scales. The southern localities usually had 6 pairs but towards the north there are only 4 or even 3. In Taiwan there were 8-12 pairs, while stems with scales be correlation between the many are more angular. There appears to no size of the flowers and the number of bracts. At some places flowers can be very small, but the plants have many pairs of bracts. kanehirae Intermediary forms between Mitrastema and M. yamamotoi occur in the same area (Yamamoto, 1925; Watanabe, 1934d), while Van Royen (1963) reported that the differences between M. kanehirae, M. kawa-sasakii, and M. yamamotoi were foundto exist in the abundant New Guinea material, which undoubtedly represented one species only. whole cline in size and numberof of scales be Overall, a pairs seems to present, and M. kanehiraeappears to be only an extreme form of M. yamamotoi. the show in which is due the In Japan plants a seasonality flowering, to presence of a dry season, in which they flower. Yamamoto distinguished a spring form (M. yamamotoi) and an autumn form ( M. kanehirae). Palm (1934) observed that in North Sumatra, where a dry season is absent, flowering occurs all around the year (see Matsuda, 1947, for dataon mean temperature and rainfall in Japan, Taiwan, and New Guinea Sumatra). In flowering seems to be seasonal, too (Van Royen, 1963). One cause of the apparent variability is possibly due to edaphic and climatic fac- tors. The depth of the roots of the hosts (Matsuda, 1947), how much litter is formed in the of the forests, mean temperature, start dry season, moisture, day length, etc. will doubt influence the host and the of its no development parasite. As some of these factors are linked with latitude, the clines couldbe explained. Matsuda (1947) suggested that the various forms are specific to their hosts. This seems an incorrect assumption, see for instance the observations by Rao & Balakrishnan (1972) on the where 4 different of Meghalaya population species 4 different families were being parasitised: Castanopsis tribuloides A. DC. (Fagaceae), Elaeocarpus lancifolius Roxb. (Elaeocarpaceae), Engelhardtia spicata Blume (Juglandaceae), and Vernonia volka- meriifolia DC. (Compositae). that has been able Now WM to study more material than any previous student of the genus, we have come to the conclusion that the Southeast Asian specimens must be regarded as representatives of a single, variable species. This might seem to some crude of but there a act lumping, are too many overlaps, gradual transitions, and too in the characters. There be narrow gaps are, as might expected, local forms, or eco- types, and in due course some of these might evolve further, if they have not been eradicated before, but at present, without looking at the field label, it is often im- with where collection and possible to say certainty a particular came from, so to what 'species' it would belong. 224 BLUMEA Vol. 38, No. 1, 1993 An example is the treatment ofthe Sumatrapopulation. Palm (1934) consideredit to be conspecific with Mitrastemayamamotoi, Hayata (in litt. to Palm) identifiedit with M. kawasasakii, but Nakai (1941a, b) based a new species on it, M. sumatranum. The Yunnan form (Li, 1975) fits in well as a range extension from Cambodia and Vietnam (Hansen, 1972, 1973). Surprisingly, in 1924 a representative of the genus was discovered in the oak for- ests of Chiapas, Mexico. It was described as a distinct species, M. matudae Y. Yama- moto (1936, ‘matudai’), because of its much larger size, broadly triangular scales, and triangular fruits, and its conical, more or less pyramidal stigma. Matsuda (1947) gave a more extensive description, based on living material. At first sight this amphi- distribution but there few similar of Pacific seems very strange, are quite a ones typi- cal tropical genera. The explanation for such a curious disjunction remains unknown (see Van Steenis, 1962, for a list more than 20 genera).
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